MYB转录因子在调控植物响应逆境胁迫中的作用

doi:10.13560/j.cnki.biotech.bull.1985.2023-1186

摘要/Abstract

摘要：

MYB作为植物中最大的多功能转录因子（transcription factors, TFs）家族之一，在基因转录水平上广泛地参与调控植物生长发育、激素信号转导及逆境胁迫应答等过程。该类转录因子N端含有典型的MYB结构域，根据MYB结构域中R重复序列的数量分为不同的亚组；而C端结构域差异较大，因此功能上具有多样性。大量研究表明，在受到外界环境信号的激活后，MYB可单独或通过和其他蛋白互作后，与下游靶基因启动子区域的顺式作用元件MYBCORE和AC-box结合，参与调控下游胁迫应答相关基因的表达，从而调节植物对逆境胁迫的耐受性。另外，MYB也通过参与脱落酸（abscisic acid, ABA）、油菜素内酯（brassinolide, BR）、茉莉酸（jasmonic acid, JA）和活性氧（reactive oxygen species, ROS）等信号通路的方式，对非生物胁迫以及生物胁迫做出应答反应。论文对植物MYB家族的结构与分类及其作用方式进行了归纳，重点对植物MYB参与调控响应盐、干旱、极端温度、营养亏缺、重金属以及病原菌等非生物和生物逆境胁迫的作用机制进行了综述，并对未来重点研究方向提出了展望，为今后农作物的抗逆性遗传改良和生物育种提供优异基因资源和理论支持。

关键词: MYB转录因子, 转录调控, DNA结合结构域, 非生物胁迫, 生物胁迫, 抗逆性, 基因表达, 蛋白互作

Abstract:

MYB is one of the biggest multifunctional transcription factors（TFs）families in plants. It is widely involved in regulating multiple process at the transcription level, including plant growth and development, signal transduction of various phyhormones and response to abiotic and biotic stresses. The N-terminal of MYB contained a typical MYB domain, which is divided into different subgroups according to the number of R repeats in the domain. However, the C-terminal domain is very different, thus it is functionally diverse. Many studies have been shown that MYB can bind to the downstream target gene promoter cis-acting elements MYBCORE and AC-box alone or through interaction with other proteins after being activated by environmental signals, and participate in regulating the expressions of downstream target genes, thereby regulating plant tolerances to stresses. Additionally, MYB responded to abiotic and biotic stresses by regulating signaling pathways such as abscisic acid（ABA）, brassinolide（BR）, jasmonic acid（JA）and reactive oxygen species（ROS）. In this review, the structure, classification, and action type of the MYB families in plants were summarized, and the regulatory mechanisms of MYB in the response to abiotic and biotic stresses such as salt, drought, extreme temperature, nutrient deficiency, heavy metals, and pathogenic bacteria, were mainly reviewed. The key direction of future studies were also prospected. This paper may provide genetic resources and theoretical support for genetic improvement of stress resistance of crops and biological breeding in the future.

Key words: MYB transcription factors, transcriptional regulation, DNA-binding domain, abiotic stress, biotic stress, resistance to stress, gene expression, protein interaction

胡雅丹, 伍国强, 刘晨, 魏明. MYB转录因子在调控植物响应逆境胁迫中的作用[J]. 生物技术通报, 2024, 40(6): 5-22.

HU Ya-dan, WU Guo-qiang, LIU Chen, WEI Ming. Roles of MYB Transcription Factor in Regulating the Responses of Plants to Stress[J]. Biotechnology Bulletin, 2024, 40(6): 5-22.

图/表 5

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物种 Species	基因名称 Gene name	基因亚家族 Gene subfamily	靶基因及位点 Target genes and sites	功能 Function	胁迫响应 Stress responses	参考文献 Reference
拟南芥 Arabidopsis thaliana	AtMYB25	R2R3-MYB	DREB2C, RD29a, SLAH1, JAZ10	激活下游胁迫应答基因	盐分、渗透性、脱落酸	[43]
	AtMYB37	R2R3-MYB	ABF2/3, COR15A, RD29a, RD22, PSII/I	提高PSII活性，调节能量耗散比例	盐分、脱落酸、干旱	[44]
	AtMYBS1	MYB-related	MAX1	负向调控独脚金素内酯途径	高温（-）	[45]
	AtMYB74	R2R3-MYB	ERF53, NIG1, HSFA6a, MYB47, MYB90, MYB102	受生长素前体IAM激活后调控下游靶基因	渗透、高温	[46]
	AtMYB12	R2R3-MYB	Flavonoid biosynthesis genes, ZEP, NCED, ABA2, AAO, P5CS, P5CR, LEA, SOD, CAT, POD	提高胁迫条件下植物中黄酮类化合物含量	盐分、干旱、高温、紫外线	[47]
	AtMYB71	R2R3-MYB	ABA response genes	调节植物ABA应答	脱落酸	[48]
	AtMYB94/96	R2R3-MYB	KCS1/2/6, KCR1, CER1/3, WSD1	促进植物表皮蜡质生物合成	干旱、强光、紫外线	[49]
花生 Arachis hypogaea	AhMYB30	MYB-related	KIN1, COR15a, RD29A, ABI2	上调参与DREB/CBF和ABA信号途径的下游胁迫相关基因表达	低温、盐分	[50]
白菜型油菜 Brassica campestris	BcMYB111	R2R3-MYB	F3H, FLS1	被CBF转录激活后增强黄酮类化合物生物合成	低温	[51]
水稻 Oryza sativa	OsMYBR57	MYB-related	OsbZIPs transcription is regulated after interaction with OsHB22	与HB22互作后激活转录因子bZIP	干旱	[52]
	OsMYB-R1	MYB-related	SOD, CAT, GPX, LEA, ABRE	激活下游胁迫相关基因	干旱、铬元素、水杨酸、脱落酸、茉莉酸	[53]
	OsFLP	R2R3-MYB	NAC1/6, DST, peroxidase 24 precursor	激活下游相关转录因子	脱落酸、干旱、盐分	[54]
陆地棉 Gossypium hirsutum	GhMYB36	R2R3-MYB	PR1	激活下游胁迫相关基因	干旱、黄萎病	[55]
	GhMYB102	R2R3-MYB	NCED1, ZAT10	参与调控ABA生物合成及干旱应答基因表达	干旱	[56]
二色补血草 Limonium bicolor	LbMYB48	MYB-related	CPC-like, DIS3, SOSs, RLKs, GSTs	调控表皮发育相关及盐胁迫相关基因表达	盐分	[57]
	LbTRY	MYB-related	GL3, ZFP5, RHD6, RSL1, LRL2/3, SOS1/2/3, P5CS	上调GL3/ZFP5表达后与表达产物竞争性结合，改变转基因植株表皮细胞分化方向，使得根毛的发育增强，吸收更多的Na⁺	盐分（-）	[58]
山核桃 Carya cathayensis	CcMYB12	R2R3-MYB	C4H, CHI, F3H, ANR, ANS, DFR	参与花青素合成途径	盐分、干旱、酸	[59]
橡胶树 Hevea brasiliensis	HbMYB44	R2R3-MYB	Homologous genes and interacting protein-encoding genes	激活下游相关基因	盐分、渗透、干旱、脱落酸、茉莉酸甲酯、赤霉素、水杨酸	[60]
甘薯 Ipomoea batatas	IbMYB308	R2R3-MYB	SOD, POD, APX, P5CS	激活下游胁迫相关基因	盐分	[61]
	IbMYB73	R2R3-MYB	NCED3, ABA2, AAO3, ABI2, SnRK2.3, DREB1D, RD22, RD26, GER5	激活ABA依赖型的不定根生长及胁迫耐受性的负调控因子转录表达	脱落酸（-）、盐分（-）、干旱（-）	[62]
灰绿藜 Chenopodium glaucum	CgMYB1	R2R3-MYB	NHX1, HAK5, SOS1, P5CS2, SOD, POD1, CBF1, COR15, COR47, bHLH001	提高转基因植物生理机能及胁迫相关基因表达	盐分、低温	[63]
豌豆 Pisum sativum	PsFLP	R2R3-MYB	CYCA2;3, CDKA;1, AAO3, NCED3, SnRK2.3	调控气孔形成及ABA合成与信号转导基因	脱落酸、干旱	[64]
84K杨 Populus alba × Populus glandulosa	PagMYB205	R2R3-MYB	POD, SOD, CAT and root vitality related genes	负调节抗氧化酶活性及根活力	盐分（-）	[65]
	PagMYB151	R2R3-MYB	Proline biosynthesis genes	与共表达转录因子共同改变根结构，促进脯氨酸积累及减少MDA含量	盐分	[66]
葡萄 V. labrusca×V. riparia	VhMYB2	R2R3-MYB	SOS1/2/3, NHX1, SnRK2.6, NCED3, P5CS1, CAT1	激活下游相关基因	高盐、干旱	[67]
辣椒 Capsicum annuum	CaDIM1	MYB-related	OSR1, RAB18, NCED3 and stress-responsive genes	诱导胁迫/ABA相关基因表达	脱落酸、干旱	[68]
荞麦 Fagopyrum tataricum	FtMYB11	R2R3-MYB	CBF1, DREB2A, RD20, ABA3, NCED3, C4H, 4CL, F3H, ANS, DFR	调节ABA信号途径、干旱及类黄酮生物合成相关基因表达水平	脱落酸、干旱（-）、盐分（-）	[69]
芹菜 Apium graveolens	AgMYB5	R2R3-MYB	CRTISO, LCYB, ABA1/2, NCED6, AAO3, ERD1, RD22, P5CS1, RD29	增强了β-胡萝卜素生物合成，随后诱导ABA合成	氧化损伤、干旱	[70]
梁 Setaria italica	SiMYB16	MYB-related	CSE, FAR1, CYP87A3, 4CL1, PAL, F5H, COMT, NCED3	调控植物木质素、类黄酮及木栓质生物合成	盐分	[71]
木薯 Manihot esculenta	MeMYB60	R2R3-MYB	CAT1/2	参与保卫细胞中活性氧稳态并影响气孔运动	脱落酸、干旱	[72]
葡萄 Vitis amurensis	VaMYB14	R2R3-MYB	ABA signaling genes, CORs, LTPs, CAT, POD	参与激活ABA信号组分和CBF-COR非依赖的LTP3表达	盐分、低温、干旱	[73]
番茄 Solanum lycopersicum	SlMYB41	R2R3-MYB	SlHSP90.3	维持热胁迫下的活性氧稳态	高温	[74]

物种 Species	基因名称 Gene name	基因亚家族 Gene subfamily	靶基因及位点 Target genes and sites	功能 Function	胁迫响应 Stress responses	参考文献 Reference
拟南芥 Arabidopsis thaliana	AtMYB25	R2R3-MYB	DREB2C, RD29a, SLAH1, JAZ10	激活下游胁迫应答基因	盐分、渗透性、脱落酸	[43]
	AtMYB37	R2R3-MYB	ABF2/3, COR15A, RD29a, RD22, PSII/I	提高PSII活性，调节能量耗散比例	盐分、脱落酸、干旱	[44]
	AtMYBS1	MYB-related	MAX1	负向调控独脚金素内酯途径	高温（-）	[45]
	AtMYB74	R2R3-MYB	ERF53, NIG1, HSFA6a, MYB47, MYB90, MYB102	受生长素前体IAM激活后调控下游靶基因	渗透、高温	[46]
	AtMYB12	R2R3-MYB	Flavonoid biosynthesis genes, ZEP, NCED, ABA2, AAO, P5CS, P5CR, LEA, SOD, CAT, POD	提高胁迫条件下植物中黄酮类化合物含量	盐分、干旱、高温、紫外线	[47]
	AtMYB71	R2R3-MYB	ABA response genes	调节植物ABA应答	脱落酸	[48]
	AtMYB94/96	R2R3-MYB	KCS1/2/6, KCR1, CER1/3, WSD1	促进植物表皮蜡质生物合成	干旱、强光、紫外线	[49]
花生 Arachis hypogaea	AhMYB30	MYB-related	KIN1, COR15a, RD29A, ABI2	上调参与DREB/CBF和ABA信号途径的下游胁迫相关基因表达	低温、盐分	[50]
白菜型油菜 Brassica campestris	BcMYB111	R2R3-MYB	F3H, FLS1	被CBF转录激活后增强黄酮类化合物生物合成	低温	[51]
水稻 Oryza sativa	OsMYBR57	MYB-related	OsbZIPs transcription is regulated after interaction with OsHB22	与HB22互作后激活转录因子bZIP	干旱	[52]
	OsMYB-R1	MYB-related	SOD, CAT, GPX, LEA, ABRE	激活下游胁迫相关基因	干旱、铬元素、水杨酸、脱落酸、茉莉酸	[53]
	OsFLP	R2R3-MYB	NAC1/6, DST, peroxidase 24 precursor	激活下游相关转录因子	脱落酸、干旱、盐分	[54]
陆地棉 Gossypium hirsutum	GhMYB36	R2R3-MYB	PR1	激活下游胁迫相关基因	干旱、黄萎病	[55]
	GhMYB102	R2R3-MYB	NCED1, ZAT10	参与调控ABA生物合成及干旱应答基因表达	干旱	[56]
二色补血草 Limonium bicolor	LbMYB48	MYB-related	CPC-like, DIS3, SOSs, RLKs, GSTs	调控表皮发育相关及盐胁迫相关基因表达	盐分	[57]
	LbTRY	MYB-related	GL3, ZFP5, RHD6, RSL1, LRL2/3, SOS1/2/3, P5CS	上调GL3/ZFP5表达后与表达产物竞争性结合，改变转基因植株表皮细胞分化方向，使得根毛的发育增强，吸收更多的Na⁺	盐分（-）	[58]
山核桃 Carya cathayensis	CcMYB12	R2R3-MYB	C4H, CHI, F3H, ANR, ANS, DFR	参与花青素合成途径	盐分、干旱、酸	[59]
橡胶树 Hevea brasiliensis	HbMYB44	R2R3-MYB	Homologous genes and interacting protein-encoding genes	激活下游相关基因	盐分、渗透、干旱、脱落酸、茉莉酸甲酯、赤霉素、水杨酸	[60]
甘薯 Ipomoea batatas	IbMYB308	R2R3-MYB	SOD, POD, APX, P5CS	激活下游胁迫相关基因	盐分	[61]
	IbMYB73	R2R3-MYB	NCED3, ABA2, AAO3, ABI2, SnRK2.3, DREB1D, RD22, RD26, GER5	激活ABA依赖型的不定根生长及胁迫耐受性的负调控因子转录表达	脱落酸（-）、盐分（-）、干旱（-）	[62]
灰绿藜 Chenopodium glaucum	CgMYB1	R2R3-MYB	NHX1, HAK5, SOS1, P5CS2, SOD, POD1, CBF1, COR15, COR47, bHLH001	提高转基因植物生理机能及胁迫相关基因表达	盐分、低温	[63]
豌豆 Pisum sativum	PsFLP	R2R3-MYB	CYCA2;3, CDKA;1, AAO3, NCED3, SnRK2.3	调控气孔形成及ABA合成与信号转导基因	脱落酸、干旱	[64]
84K杨 Populus alba × Populus glandulosa	PagMYB205	R2R3-MYB	POD, SOD, CAT and root vitality related genes	负调节抗氧化酶活性及根活力	盐分（-）	[65]
	PagMYB151	R2R3-MYB	Proline biosynthesis genes	与共表达转录因子共同改变根结构，促进脯氨酸积累及减少MDA含量	盐分	[66]
葡萄 V. labrusca×V. riparia	VhMYB2	R2R3-MYB	SOS1/2/3, NHX1, SnRK2.6, NCED3, P5CS1, CAT1	激活下游相关基因	高盐、干旱	[67]
辣椒 Capsicum annuum	CaDIM1	MYB-related	OSR1, RAB18, NCED3 and stress-responsive genes	诱导胁迫/ABA相关基因表达	脱落酸、干旱	[68]
荞麦 Fagopyrum tataricum	FtMYB11	R2R3-MYB	CBF1, DREB2A, RD20, ABA3, NCED3, C4H, 4CL, F3H, ANS, DFR	调节ABA信号途径、干旱及类黄酮生物合成相关基因表达水平	脱落酸、干旱（-）、盐分（-）	[69]
芹菜 Apium graveolens	AgMYB5	R2R3-MYB	CRTISO, LCYB, ABA1/2, NCED6, AAO3, ERD1, RD22, P5CS1, RD29	增强了β-胡萝卜素生物合成，随后诱导ABA合成	氧化损伤、干旱	[70]
梁 Setaria italica	SiMYB16	MYB-related	CSE, FAR1, CYP87A3, 4CL1, PAL, F5H, COMT, NCED3	调控植物木质素、类黄酮及木栓质生物合成	盐分	[71]
木薯 Manihot esculenta	MeMYB60	R2R3-MYB	CAT1/2	参与保卫细胞中活性氧稳态并影响气孔运动	脱落酸、干旱	[72]
葡萄 Vitis amurensis	VaMYB14	R2R3-MYB	ABA signaling genes, CORs, LTPs, CAT, POD	参与激活ABA信号组分和CBF-COR非依赖的LTP3表达	盐分、低温、干旱	[73]
番茄 Solanum lycopersicum	SlMYB41	R2R3-MYB	SlHSP90.3	维持热胁迫下的活性氧稳态	高温	[74]